“Mail Online continues to focus on attracting traffic directly to its homepages on desktop and mobile or its apps,” said Daily Mail & General Trust (DMGT), the parent company of the Mail titles and the Metro freesheet. “Indirect traffic, notably via social media and search platforms, has reduced.”

Despite the audience slump, Mail Online said the total minutes spent on the site increased to a daily average of 145m minutes, with more than three-quarters of that from direct traffic. Mail Online made £122m in advertising revenues in the year to the end of September, a 3% year-on-year rise.

The company said the print ad decline “reflects the continued structural and competitive challenges facing the UK national newspaper advertising market”.

Paul Zwillenberg, the chief executive of DMGT, said: “MailOnline continues to perform well and has reached an important milestone with digital advertising revenue now exceeding the Mail’s print advertising revenues.”

Overall there was a 7% decline in total revenues, including copy sales income, at the Mail print titles to £424m. Revenues at the Mail businesses in totality – the print titles and Mail Online combined – fell 5% to £546m.

Metro, which was put up for sale this year, reported a 4% increase in ad revenues to £71m for the year.